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COMPOSITION OF THE GRAM-POSITIVE PLASMA MEMBRANE Protein ( 65%; large range [>200] distinct protein species present) Lipid ( 25%; mainly phospholipid; 3-4 major types present) Lipoteichoic acid (5-10%) MAJOR PHOSPHOLIPID HEADGROUP (X) SPECIES IN BACTERIA -X PHOSPHOLIPID -H Phosphatidic acid (PA) -CH2-CH2-NH2 Phosphatidyl ethanolamine (PE) -CH2-CH-NH2 CO2H Phosphatidyl serine (PS) -CH2-CH-CH2 OH OH Phosphatidyl glycerol (PG) O -CH2-CH-CH2-O-P-OOH O CH2 O CH-O-C-R1 CH2-O-C-R2 O Diphosphatidyl glycerol/Cardiolipin (diPG/CL) (R1 & R2 = long [ C16] acyl chains) FUNCTIONS OF GRAM-POSITIVE PLASMA MEMBRANE 1. 3. 4. 6. 8. 9. Highly selective permeability barrier. 2. Concentrative solute transport. Oxidative phosphorylation and electron transport. Biosynthesis of peptidoglycan. 5. Biosynthesis of teichoic/teichuronic acids. Biosynthesis of lipoteichoic acids. 7. Biosynthesis of capsular polysaccharides. Biosynthesis of phospholipids. 8. Translocation and processing of exported proteins Chemosensing and transmembrane signaling. 10. Chromosome anchoring. COMPOSITION & FUNCTIONS OF GRAM-VE PLASMA (INNER) MEMBRANE Protein and phospholipid composition generally similar to that of Gram+ves BUT Lipoteichoic acids NOT found Functionally similar to Gram+ve plasma membrane BUT No biosynthesis of wall teichoic acids or membrane lipoteichioc acids. Biosynthesis of lipopolysaccharides and membrane-derived oligosaccharides Additional transport system (osmotic shock sensitive [periplasmic] transport system) COMPOSITION OF THE (GRAM-VE) PERIPLASM Water soluble proteins Low-molecular-weight carbohydrate “membrane derived oligosaccharides (MDO)” (Peptidoglycan layer) FUNCTIONS OF THE (GRAM-VE) PERIPLASM 1. 2. 3. 4. 5. 6. Nutrient degradation (phosphatases, carboxypeptidases) Nutrient transport (specific periplasmic binding proteins) Electron transport (denitrifying and chemolithotrophic bateria) Elimination of foreign DNA (restriction endonucleases) Modification of potentially toxic compounds (-lactamase; aminoglycoside 3’ phosphotransferase) Maintain iso-osmolarity across the plasma membrane (MDO) [P-glycerol]1-4 EtNP-[Glucose]8-10 O-succinate Fig. 17. Membrane-derived oligosaccharide (MDO). EtNP = phosphoethanolamine. Note that glycerol phosphate and succinate both carry –ve charges. COMPOSITION OF THE GRAM-VE PEPTIDOGLYCAN LAYER Peptidoglycan (relatively thin 2nm) Covalently bound lipoprotein O R1-C-O-CH2 R2-C-O-CH diglyceride O CH2 S thioether linkage CH2 O R3-C-N-C-C-{56 amino acids}-Lys(58)-Diaminopimelic acid (of peptidoglycan) H HO Fig. 18. The Braun lipoprotein of E. coli. The N-terminal cysteine (Cys1) residue is modified by the attachment of amide-linked fatty acid (R3-CO) and thioether-linked diglyceride. The three fatty acyl groups interdigitate into the inner leaflet of the outer membrane whilst the C-terminal lysine (Lys58) is amide-linked to the DAP of peptidoglycan. Thus, the Braun lipoprotein effectively anchors the outer membrane to the underlying murein (see Fig. 19 & 20).